{"database":"Pride","file_versions":[{"headers":{"Content-Type":["application/json"]},"body":{"files":{"Xlsx":["ftp://ftp.pride.ebi.ac.uk/pride/data/archive/2026/07/PXD072461/Additives_Rapigest02.raw_20260425_Byonic.xlsx","ftp://ftp.pride.ebi.ac.uk/pride/data/archive/2026/07/PXD072461/Additives_Rapigest01_CM830_3.raw_20260425_Byonic.xlsx","ftp://ftp.pride.ebi.ac.uk/pride/data/archive/2026/07/PXD072461/Additives_none_02.raw_20260425_Byonic.xlsx","ftp://ftp.pride.ebi.ac.uk/pride/data/archive/2026/07/PXD072461/VKOR_trypsin_02.raw_20260425_Byonic.xlsx","ftp://ftp.pride.ebi.ac.uk/pride/data/archive/2026/07/PXD072461/VKOR_Pepsin01_CM830_8.raw_20260425_Byonic.xlsx","ftp://ftp.pride.ebi.ac.uk/pride/data/archive/2026/07/PXD072461/VKOR_trypsin_01.raw_20260425_Byonic.xlsx","ftp://ftp.pride.ebi.ac.uk/pride/data/archive/2026/07/PXD072461/Additives_Urea_02.raw_20260425_Byonic.xlsx","ftp://ftp.pride.ebi.ac.uk/pride/data/archive/2026/07/PXD072461/VKOR_Thermolysin02_CM828_10RR.raw_20260425_Byonic.xlsx","ftp://ftp.pride.ebi.ac.uk/pride/data/archive/2026/07/PXD072461/Additives_ProteaseMAX02_CM830_2.raw_20260425_Byonic.xlsx","ftp://ftp.pride.ebi.ac.uk/pride/data/archive/2026/07/PXD072461/Additives_none_01.raw_20260425_Byonic.xlsx","ftp://ftp.pride.ebi.ac.uk/pride/data/archive/2026/07/PXD072461/Additives_TFE01_CM831_5R.raw_20260425_Byonic.xlsx","ftp://ftp.pride.ebi.ac.uk/pride/data/archive/2026/07/PXD072461/Additives_ProteaseMAX01_CM830_2R.raw_20260425_Byonic.xlsx","ftp://ftp.pride.ebi.ac.uk/pride/data/archive/2026/07/PXD072461/Additives_TFE02_CM831_5.raw_20260425_Byonic.xlsx","ftp://ftp.pride.ebi.ac.uk/pride/data/archive/2026/07/PXD072461/VKOR_Pepsin02.raw_20260425_Byonic.xlsx","ftp://ftp.pride.ebi.ac.uk/pride/data/archive/2026/07/PXD072461/Additives_Urea_01_CM828_2R_NORMAL.raw_20260503_Byonic.xlsx","ftp://ftp.pride.ebi.ac.uk/pride/data/archive/2026/07/PXD072461/VKOR_Thermolysin01_20260425_Byonic.xlsx"],"Raw":["ftp://ftp.pride.ebi.ac.uk/pride/data/archive/2026/07/PXD072461/Additives_TFE02.raw","ftp://ftp.pride.ebi.ac.uk/pride/data/archive/2026/07/PXD072461/VKOR_Therrmolysin_01.raw","ftp://ftp.pride.ebi.ac.uk/pride/data/archive/2026/07/PXD072461/Additives_Rapigest_02.raw","ftp://ftp.pride.ebi.ac.uk/pride/data/archive/2026/07/PXD072461/Additives_none_02.raw","ftp://ftp.pride.ebi.ac.uk/pride/data/archive/2026/07/PXD072461/VKOR_Pepsin_02.raw","ftp://ftp.pride.ebi.ac.uk/pride/data/archive/2026/07/PXD072461/Additives_Rapigest01.raw","ftp://ftp.pride.ebi.ac.uk/pride/data/archive/2026/07/PXD072461/VKOR_Chymotrypsin_02.raw","ftp://ftp.pride.ebi.ac.uk/pride/data/archive/2026/07/PXD072461/Vkor_Thermolysin02.raw","ftp://ftp.pride.ebi.ac.uk/pride/data/archive/2026/07/PXD072461/Additives_Urea01.raw","ftp://ftp.pride.ebi.ac.uk/pride/data/archive/2026/07/PXD072461/VKOR_Chymotrypsin_01.raw","ftp://ftp.pride.ebi.ac.uk/pride/data/archive/2026/07/PXD072461/Additives_none_01.raw","ftp://ftp.pride.ebi.ac.uk/pride/data/archive/2026/07/PXD072461/Additives_ProteaseMAX01.raw","ftp://ftp.pride.ebi.ac.uk/pride/data/archive/2026/07/PXD072461/Vkor_Pepsin01.raw","ftp://ftp.pride.ebi.ac.uk/pride/data/archive/2026/07/PXD072461/Additives_TFE01.raw","ftp://ftp.pride.ebi.ac.uk/pride/data/archive/2026/07/PXD072461/VKOR_trypsin_02.raw","ftp://ftp.pride.ebi.ac.uk/pride/data/archive/2026/07/PXD072461/Additives_urea_02.raw","ftp://ftp.pride.ebi.ac.uk/pride/data/archive/2026/07/PXD072461/Additives_ProteaseMAX02.raw","ftp://ftp.pride.ebi.ac.uk/pride/data/archive/2026/07/PXD072461/VKOR_trypsin_01.raw"],"Fasta":["ftp://ftp.pride.ebi.ac.uk/pride/data/archive/2026/07/PXD072461/selected_proteins.fasta","ftp://ftp.pride.ebi.ac.uk/pride/data/archive/2026/07/PXD072461/VKOR_Ecoli.fasta"]},"type":"primary"},"statusCode":"OK","statusCodeValue":200}],"scores":null,"additional":{"labhead_mail":["mgross@wustl.edu"],"submitter":["Ming Cheng"],"technology_type":["Mass Spectrometry","Bottom-up proteomics"],"software":["Not available"],"submitter_keywords":["filter-aided sample preparation (fasp)","Membrane protein digestion","Membrane protein","protein footprinting","mass spectrometry","Membrane protein; vitamin k epoxide reductase (vkor); mass spectrometry; protein footprinting; filter-aided sample preparation (fasp)","vitamin k epoxide reductase (vkor)"],"full_dataset_link":["https://www.ebi.ac.uk/pride/archive/projects/PXD072461"],"sample_protocol":["The FASP protocol was modified on the basis of previous literature [52] (the workflow for FASP is described in Fig. 1 in the main text). Specifically, ultrafiltration units contain-ing 50 µL VKOR (10 µM) were added to 200 μL of denatured solution containing 8 M urea in 0.1 M ammonium bicarbonate (pH 8.0) in the ultrafiltration units and centrifuged at 10,000g for ~ 20 min until less than 10 μL of sample remains above the filter. Then, 200 μL of denatured solution was added to the ultrafiltration units, and this wash step was re-peated twice. The flow-through from the collection tube was discarded, and  100 μL 50 mM TCEP in 0.1 M ammonium bicarbonate was added to the filter and incubated at 37  °C for 30 min. Iodoacetamide (IAA, 100 μL, 100 mM) in 0.1 M ammonium bicarbonate was added, and the resulting solution was placed in the dark for 30 min. The ultrafiltra-tion units were centrifuged at 10,000 g for ~ 20 min, and 150 μL of digestion buffer (DB, 0.1 M ammonium bicarbonate) was added to the filtration units, and they were centrifuged at 10,000 g for 10 min. This step was repeated twice. At this stage, urea and excess IAA and TCEP were removed by DB wash, and 60 μL of DB and protease (enzyme to protein ratio 1:20 (w/w) )was added. For pepsin digestion, 0.1 N HCl in water was used as the digestion buffer.  The units were placed in a water bath at 37 °C overnight. Note: The incubation of ultrafiltration units in a thermomixer will result in evaporation of DB overnight. The units were kept in water bath or a humid chamber at 37 °C overnight. After digestion, the ultrafiltra-tion units were transferred to new collection tubes, and they were centrifuged 10,000 g un-til the solution completely passed the filter membrane. A volume of 100 μL of DB was added, and the ultrafiltration units were centrifuged at 10,000 g until the solution com-pletely passed the filter membrane. This step was repeated. The flow-through was col-lected, and 1 μL formic acid was added to acidify the solution. The flow-through contain-ing proteolytic peptides wasused for subsequent MS analysis. The concentration of the peptides was determined by using a Thermo Scientific™ NanoDrop™.,"],"repository":["Pride"],"quantification_method":["Not available"],"modification":[""],"data_protocol":["LC-MS/MS raw files were imported into the Byonic™ Software (Protein Metrics, San Car-los, CA, USA) for peptide mapping. Byonic™ processing of the LC-MS/MS data directly provided sequence coverage, number of unique peptides under different proteolytic condi-tions. Chosen were 20 ppm precursor mass tolerance, 60 ppm fragment mass tolerance, and CID/HCD fragmentation. Peptide identifications were conducted by searching the raw data against the VKOR sequence. The search parameters used were as follows: for trypsin, the digest mode is semispecific on K and R, and the “max missed” cleavages were four. When nonspecific proteases (i.e., chymotrypsin, thermolysin, pepsin) were used, the digest mode was set as nonspecific. Carbamidomethylation (+57.0215) of Cys was set as common modification, oxidation (+15.9945 Da) of Met, Tyr, Phe, Trp, and carbamylation (+ 43.0058 Da) of Thr, Ser and Lys was set as variable modifications."],"omics_type":["Proteomics"],"labhead":["Michael L Gross"],"instrument_platform":[""],"labhead_affiliation":["Washington University in St. Louis"],"submission_type":["PARTIAL"],"species":["Homo Sapiens (human)"],"publication":["10.3390/MEMBRANES16060215"],"submitter_mail":["chengming@simm.ac.cn"],"submitter_affiliation":["Shanghai Institute of Materia Medica Chinese Academy of Sciences"],"submitter_country":["China"],"additional_accession":[]},"is_claimable":false,"name":"Optimized Digestion Conditions for Membrane Protein Foot-printing and Mass Spectrometry Analysis","description":"Integral membrane proteins (IMPs), which constitute 50–60% of drug targets, play essen-tial roles in numerous biological processes but remain underrepresented in conventional bottom-up and in structural proteomics owing to their hydrophobicity and resistance to proteolysis. Although advances in IMPs proteomics have improved global IMPs detection, most efforts focus on proteome-scale protein identification rather than targeted structural analysis. Protein footprinting in structural proteomics, require high sequence coverage and protein digestion to peptides of suitable length for structural elucidation, necessitating optimized digestion condition for individual IMPs. Here, we report a digestion protocol tailored for structural mass spectrometry and evalu-ate its performance using a single amphipathic IMP model featuring distinct extramem-brane and transmembrane domains. We evaluated the use of various protease–additive combinations and applied filter-aided sample preparation (FASP) to remove detergents and surfactants efficiently prior to MS analysis. The optimized conditions consistently yielded >90% sequence coverage. Guided by MS retention time calibration and hydropho-bic factor simulations, we identified a “sweet spot” for transmembrane peptide detection. Notably, although cleavable surfactants can enhance proteome-wide coverage, our results show that they are not essential for single protein studies as in structural proteomics. In-stead, detergent removal, protease selection, and generation of suitably sized peptides are critical for enabling reliable bottom-up structural analysis of IMPs. The protocol developed here provides a practical framework for optimizing digestion conditions in IMPs charac-terization.","dates":{"publication":"2026-07-02","submission":"2025-12-28"},"accession":"PXD072461","cross_references":{"TAXONOMY":["NEWT:330879","NEWT:377960","NEWT:1131","NEWT:35795","NEWT:2042546","NEWT:259447","NEWT:295546","NCBITaxon:2719036","NEWT:1334565","NEWT:625351","NEWT:258594","NCBITaxon:1280","NEWT:112503","NEWT:1129","NEWT:309807","NEWT:59753","NEWT:89184","NEWT:309800","NEWT:281395","NEWT:1211601","NEWT:876138","NEWT:44271","NEWT:193516","NEWT:2315688","NEWT:1073882","NEWT:6819","NEWT:475174","NEWT:111205","NEWT:428386","NEWT:1117","NEWT:498257","NEWT:10036","NEWT:32264","NEWT:12215","NEWT:1590","NEWT:10034","NEWT:33114","NEWT:1596","NEWT:661410","NEWT:638632","NEWT:224326","NEWT:376619","NCBITaxon:79857","NEWT:1096976","NEWT:3765","NEWT:3767","NEWT:1589","NEWT:135622","NEWT:4615","NEWT:672127","NEWT:35786","NEWT:1580","NEWT:399784","NEWT:96731","NEWT:656064","NEWT:1582","NEWT:3760","NEWT:1584","NEWT:36630","NEWT:1585","NEWT:383379","NEWT:418106","NCBITaxon:12721","NEWT:10029","NEWT:913645","NEWT:2492962","NEWT:641809","NEWT:1392488","NEWT:38815","NEWT:317447","NEWT:4688","NEWT:111225","NEWT:7719","NEWT:5530","NEWT:868565","NEWT:135674","NEWT:79329","NEWT:30069","NEWT:12637","NEWT:59729","NEWT:2164133","NEWT:295105","NEWT:108061","NEWT:60711","NEWT:224308","NEWT:3347","NEWT:49928","NEWT:3349","NEWT:160621","NEWT:212790","NEWT:110368","NEWT:55350","NEWT:1310161","NEWT:77133","NEWT:145481","NEWT:1310165","NEWT:29058","NCBITaxon:79824","NEWT:1912919","NEWT:410658","NEWT:5516","NEWT:44688","NEWT:44689","NEWT:498211","NEWT:345076","NEWT:347256","NEWT:5518","NEWT:398007","NEWT:1527468","NEWT:498217","NEWT:498216","NEWT:11320","NEWT:246196","NEWT:246197","NEWT:145458","NEWT:1310173","NEWT:44685","NEWT:161934","NEWT:156587","NEWT:1148","NEWT:5508","NEWT:3329","NEWT:419481","NEWT:5507","NEWT:260499","NEWT:410661","NEWT:1140","NCBITaxon:2157","NEWT:2066563","NEWT:1143","NEWT:1287689","NEWT:1094343","NEWT:1462472","NEWT:1336795","NEWT:644042","NEWT:150808","NEWT:1182590","NEWT:3712","NEWT:3711","NEWT:3714","NCBITaxon:37162","NEWT:270643","NEWT:2065263","NEWT:177437","NEWT:717966","NEWT:10891","NEWT:10418","NEWT:118698","NEWT:1616117","NEWT:118696","NEWT:34865","NEWT:52283","NEWT:28174","NEWT:284812","NEWT:8175","NEWT:43330","NEWT:1603293","NEWT:44664","NEWT:43346","NEWT:3702","NEWT:1245466","NEWT:3704","NEWT:244366","NEWT:1246791","NEWT:176597","NEWT:118694","NEWT:2850","NEWT:118691","NEWT:34871","NEWT:33548","NEWT:52299","NEWT:8165","NEWT:96794","NEWT:3708","NEWT:332648","NEWT:44670","NEWT:926571","NEWT:536231","NEWT:376219","NEWT:219813","NEWT:1510","NEWT:460519","NEWT:1515","NEWT:572307","NEWT:1432138","NEWT:1424507","NEWT:1194599","NEWT:272844","NEWT:1348799","NEWT:33995","NEWT:55784","NEWT:1303443","NEWT:9483","NEWT:485","NEWT:56636","NEWT:2709072","NEWT:2499525","NEWT:2853422","NEWT:487","NEWT:1538553","NEWT:1679718","NEWT:480","NEWT:67767","NEWT:45972","NEWT:46835","NEWT:109757","NEWT:582580","NEWT:1185650","NEWT:216778","NEWT:294607","NEWT:1502","NEWT:128017","NEWT:2836","NEWT:376686","NEWT:237610","NEWT:95486","EFO:0001352","NEWT:119562","NEWT:1883446","NEWT:1233435","NEWT:118223","NEWT:11757","NEWT:987059","NEWT:109760","NEWT:29031","NEWT:317047","NEWT:1509","NEWT:491","NEWT:460523","NEWT:45954","NEWT:235443","NEWT:1579","NEWT:1757312","NEWT:85694","NEWT:109786","NEWT:108458","NEWT:5936","NEWT:321967","NEWT:320637","NEWT:1078283","NEWT:3750","NEWT:983964","NEWT:11706","NEWT:32644","NEWT:527796","NEWT:499175","NEWT:109779","NEWT:3745","NEWT:1715989","NCBITaxon:4751","NEWT:3747","NEWT:1116234","NEWT:1255228","NEWT:410289","NEWT:191701","NEWT:373153","NEWT:1640278","NEWT:472","NEWT:940825","NEWT:29459","NEWT:529507","NEWT:1071661","NEWT:28128","NEWT:470","NEWT:5911","NEWT:748693","NCBITaxon:50557","NEWT:39251","NEWT:29491","NEWT:101841","NEWT:446","NEWT:153481","NEWT:2014887","NEWT:33952","NEWT:11723","NEWT:445","NEWT:153009","NEWT:261756","NEWT:166313","NEWT:63366","NEWT:63367","NEWT:215402","NEWT:1547","NEWT:1290454","NEWT:906914","NEWT:81231006","NEWT:28141","NEWT:9031","NEWT:27292","NEWT:1050722","NEWT:3728","NEWT:108931","NEWT:1293497","NEWT:1055524","NEWT:548","NEWT:150475","NEWT:267872","NEWT:381666","NEWT:172269","NEWT:9534","NEWT:2126348","NEWT:5180","NEWT:256737","NEWT:9541","NEWT:8694","NEWT:33936","NEWT:8692","NEWT:2903","NEWT:185579","NEWT:872227","NEWT:13076","NEWT:2587410","NEWT:207340","NEWT:1006581","NEWT:8673","NEWT:33940","NEWT:2126337","NEWT:550","NEWT:33941","NEWT:554","NEWT:451516","NEWT:552","NEWT:1325291","NEWT:42410","NEWT:36185","NEWT:1054211","NEWT:1225786","NEWT:771870","NEWT:51338","NEWT:9995","NEWT:575412","NEWT:28112","NEWT:49012","NEWT:6493","NEWT:10832","NEWT:6494","NEWT:6491","NEWT:507601","NEWT:520","NEWT:64657","NEWT:186441","NEWT:643680","NEWT:214092","NCBITaxon:6157","NEWT:13095","NEWT:162425","NEWT:104105","NEWT:216257","NEWT:9986","NEWT:8654","NEWT:8658","NEWT:1268063","NEWT:8657","NEWT:9983","NEWT:8655","NEWT:5147","NEWT:595495","NEWT:28108","NEWT: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